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(Orchidaceae) Во Флоре России the Genus Epipactis Zinn

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(Orchidaceae) Во Флоре России the Genus Epipactis Zinn Turczaninowia 21 (4): 19–34 (2018) ISSN 1560–7259 (print edition) DOI: 10.14258/turczaninowia.21.4.3 TURCZANINOWIA http://turczaninowia.asu.ru ISSN 1560–7267 (online edition) УДК 582.594.2(470+571) Род Epipactis Zinn (Orchidaceae) во флоре России А. В. Фатерыга, В. В. Фатерыга Карадагская научная станция им. Т. И. Вяземского – природный заповедник РАН, ул. Науки, 24, пгт Курортное, г. Феодосия, Республика Крым, 298188, Россия. E-mail: [email protected] Ключевые слова: Евразия, орхидеи, распространение, синонимия, таксономия. Аннотация. Род Epipactis Zinn является одним из наиболее таксономически сложных родов семейства Orchidaceae. Большинство видов рода составляют узкоареальные самоопыляющиеся таксоны, изолирован- ные друг от друга вследствие автогамии. Однако в последние годы можно наблюдать тенденцию к описанию большого числа перекрестноопыляющихся видов, ведущую к таксономической «инфляции». В настоящей пу- бликации приводится обзор рода Epipactis для территории России, который включает 15 видов и три допол- нительных подвида: E. palustris (L.) Crantz, E. thunbergii A. Gray, E. atrorubens (Hoffm.) Besser, E. condensata Boiss. ex D. P. Young (= E. viridiflora subsp. kuenkeleana Akhalk., H. Baumann, R. Lorenz et Mosul., syn. nov.), E. krymmontana Kreutz, Fateryga et Efimov, E. microphylla (Ehrh.) Sw., E. helleborine (L.) Crantz subsp. hellebori- ne, E. helleborine subsp. papillosa (Franch. et Sav.) Fateryga, stat. nov., E. helleborine subsp. tremolsii (Pau) E. Klein (= E. turcica Kreutz, syn. nov.; = E. helleborine subsp. levantina Kreutz, Óvári et Shifman, syn. nov.), E. distans Arv.- Touv., E. purpurata Sm., E. leptochila (Godfery) Godfery subsp. leptochila, E. leptochila subsp. neglecta Kümpel, E. muelleri Godfery, E. pontica Taubenheim, E. tangutica Schltr., E. euxina Fateryga, Popovich et Kreutz и E. persica (Soó) Hausskn. ex Nannf. (= E. taurica Fateryga et Kreutz, syn. nov.). Приводится ключ для определения видов и подвидов. The genus Epipactis Zinn (Orchidaceae) in the flora of Russia A. V. Fateryga, V. V. Fateryga T. I. Vyazemsky Karadag Scientific Station – Nature Reserve of RAS, Nauki str., 24, Kurortnoye, Feodosiya, Republic of Crimea, 298188, Russian Federation Keywords: distribution, Eurasia, orchids, synonymy, taxonomy. Summary. The genus Epipactis Zinn is one of the most complicated genera of the family Orchidaceae. The ma- jority of species in the genus are locally evolved self-pollinating taxa isolated from each other due to autogamy. A tendency to describe a large amount of cross-pollinating species, leading to taxonomic inflation, however, can be observed in recent years. The present paper revises the genus Epipactis within the territory of Russia. Fifteen species and three additional subspecies are recognized: E. palustris (L.) Crantz, E. thunbergii A. Gray, E. atrorubens (Hoffm.) Besser, E. condensata Boiss. ex D. P. Young (= E. viridiflora subsp. kuenkeleana Akhalk., H. Baumann, R. Lorenz et Mosul., syn. nov.), E. krymmontana Kreutz, Fateryga et Efimov,E. microphylla (Ehrh.) Sw., E. helleborine (L.) Crantz subsp. helleborine, E. helleborine subsp. papillosa (Franch. et Sav.) Fateryga, stat. nov., E. helleborine subsp. trem- olsii (Pau) E. Klein (= E. turcica Kreutz, syn. nov.; = E. helleborine subsp. levantina Kreutz, Óvári et Shifman, syn. nov.), E. distans Arv.-Touv., E. purpurata Sm., E. leptochila (Godfery) Godfery subsp. leptochila, E. leptochila subsp. neglecta Kümpel, E. muelleri Godfery, E. pontica Taubenheim, E. tangutica Schltr., E. euxina Fateryga, Popovich et Kreutz, and E. persica (Soó) Hausskn. ex Nannf. (= E. taurica Fateryga et Kreutz, syn. nov.). A key to the species and subspecies is provided. Поступило в редакцию 04.04.2018 Submitted 04.04.2018 Принято к публикации 22.11.2018 Accepted 22.11.2018 20 Фатерыга А. В., Фатерыга В. В. Род Epipactis (Orchidaceae) во флоре России Введение 2012). Такой способ опыления характерен для E. palustris (L.) Crantz из секции Arthrochilium и Род Epipactis Zinn (дремлик) относится к од- E. atrorubens (Hoffm.) Besser из секции Epipac- ному из наиболее таксономически сложных ро- tis s. str. (Nilsson, 1978; Brantjes, 1981; Jakubska- дов орхидных. В мировой флоре известно при- Busse, Kadej, 2011). При этом наиболее эф- мерно 50–70 видов этого рода, бо́льшая часть ко- фективными опылителями E. palustris служат торых произрастает в Евразии. По одному виду складчатокрылые осы (осы семейства Vespidae) Epipactis известно в Центральной и Северной (Nilsson, 1978), а для E. atrorubens характерно Африке, кроме того, два вида, из которых один также факультативное самоопыление (Burns- адвентивный, произрастают в Северной Амери- Balogh et al., 1987). Дальнейшая эволюция опы- ке (Efimov, 2004; Govaerts et al., 2005–2018; Chase лительных синдромов в секции Arthrochilium et al., 2015; Delforge, 2016). В состав рода входят шла по пути специализации к привлечению на две неравные по объему секции: Arthrochilium цветки мух-журчалок (мух семейства Syrphidae), Irmisch и Epipactis s. str. (= Euepipactis Irmisch). являющихся основными опылителями таких ви- Секция Arthrochilium включает 11–12 видов и дов, как E. thunbergii A. Gray (Sugiura, 1996) и иногда рассматривается как самостоятельный E. veratrifolia Boiss. et Hohen. (Stökl et al., 2011). род (Efimov, 2007). Однако, согласно самой при- При этом у E. veratrifolia обнаружены специали- знанной в настоящее время классификации ор- зированные олфакторные аттрактанты, привле- хидных (Chase et al., 2015), виды этой секции кающие именно этих мух. Данные аттрактанты входят в род Epipactis. Секцию Arthrochilium, в сходны с «феромонами тревоги» тлей, а самки свою очередь, подразделяют на ряды Longifoliae многих мух-журчалок откладывают яйца в коло- Nevski ex Efimov и Palustres Nevski ex Efimov нии тлей, поскольку их личинки питаются тля- (Efimov, 2004) либо на неформальные группы ми. Таким образом, E. veratrifolia привлекает на видов «E. veratrifolia» и «E. palustris» (Delforge, свои цветки мух, занятых поиском места для от- 2016). Подавляющее большинство видов рода кладки яиц (Stökl et al., 2011). Epipactis относится к секции Epipactis s. str. Ее Эволюция опылительных синдромов в сек- также подразделяют на ряды Atrorubentae Nevski ции Epipactis s. str. шла по пути выработки спец- ex Efimov и Epipactis s. str. (Efimov, 2004). Тем ифических аттрактантов для складчатокрылых не менее филогенетические исследования не ос, а затем их утраты и переходу к самоопыле- подтверждают монофилетичность этих рядов нию (Fateryga, Ivanov, 2012). Стоит отметить, (Tranchida-Lombardo et al., 2011; Zhou, Jin, 2018). что почти всем изученным перекрестноопыля- П. Дельфорж вместо рядов выделяет в составе ющимся (аллогамным) видам секции свойствен- секции Epipactis s. str. неформальные группы но и факультативное самоопыление (автогамия) видов «E. atrorubens», «E. tremolsii», «E. helle- (Burns-Balogh et al., 1987). Однако следует от- borine», «E. purpurata», «E. leptochila», «E. al- делять виды, у которых перекрестное опыление bensis» и «E. phyllanthes» (Delforge, 2016). Эти все же возможно, от видов, строение цветков ко- группы основаны на признаках опушения оси торых не позволяет насекомым выносить из них соцветия и завязей, листорасположения, а также поллинии. Последние характеризуются тем, что окраски цветоножки. Очевидно, что большин- их прилипальце утратило клейкость либо полно- ство из них вряд ли могут иметь серьезный так- стью редуцировалось; кроме того, у некоторых сономический вес. Поэтому данные группы слу- видов редуцировался и клювик, из-за чего пол- жат лишь для целей удобства составления опре- линий попадает на рыльце еще в бутоне. Среди делительных ключей. Кроме того, некоторые из перекрестноопыляющихся видов наиболее из- таких признаков (например, окраска цветонож- вестны E. helleborine (L.) Crantz и E. purpurata ки) могут быть достаточно лабильны в пределах Sm., привлекающие на цветки главным образом одного вида. рабочих особей общественных ос (Darwin, 1862; Высокое видовое разнообразие и таксономи- Judd, 1971; Jakubska-Busse, Kadej, 2011). В соста- ческая сложность рода Epipactis тесно связаны со ве нектара E. helleborine обнаружено несколько способами опыления этих растений. Первичным десятков вторичных метаболитов, часть из кото- опылительным синдромом в обеих секциях рода рых оказывают наркотическое действие на насе- следует считать перекрестное опыление цветков комых. Считается, что «пьяные» осы являются широким кругом опылителей, включающим ос, более эффективными опылителями, поскольку у пчел, мух и отчасти муравьев (Fateryga, Ivanov, них нарушена координация движений и они не Turczaninowia 21 (4): 19–34 (2018) 21 способны избавиться от поллиниев (Jakubska et «стройные»), степени раскрытия околоцветни- al., 2005). Некоторые из этих наркотических ве- ка (широко или не очень широко) или окраски ществ образуются в нектаре благодаря деятель- цветков (более или менее яркая). Многие из по- ности симбиотических дрожжевых грибов, кото- добных признаков (такие, как число генератив- рых заносят в гипохилий цветка сами же опыли- ных стеблей, волнистость листовой пластинки и тели (Ehlers, Olesen, 1997). Кроме того, известно, степень очередности/супротивности листьев на что привлекательность цветков E. helleborine для стебле) уже показали свою несостоятельность в ос обеспечивается также сходством выделяемого таксономии рода Epipactis (Jakubska-Busse et al., ими запаха со спектром летучих вторичных мета- 2012, 2017; Jakubska-Busse, Gola, 2014). Кроме болитов, выделяемых листьями растений при их того, диагнозы большинства недавно описанных повреждении
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